Melatonin can induce year-round ovarian cyclicity in red deer (Cervus elaphus)

From 17 February 1987 (Day 1) to 5 June 1988 (Day 475), 6 red deer hinds which had been in natural daylength (NL/M) and 6 hinds which had been in continuous artificial light for the previous month (CL/M) were each given melatonin (5 mg in feed) daily at 15:00 h. Six controls (C) received unsupplemented feed. From Day 1 all hinds were in natural daylight and ovarian cyclicity was assessed from plasma progesterone concentrations. Group C first went into anoestrus on 15 March 1987 (Day 27 +/- 9.2 (s.e.m], recommenced cyclicity on 23 October (Day 249 +/- 2.3) and went into anoestrus again on 2 April 1988 (Day 411 +/- 8.7). Group CL/M first went into anoestrus 31 days earlier (P less than 0.05) on 12 February (Day -4 +/- 7.8), before the start of melatonin treatment; 4 hinds then recommenced ovarian cycles 132 days earlier (P less than 0.001) on 13 June (Day 117 +/- 5.8) and continued to cycle for a longer period than did controls. Group NL/M hinds were cyclic at the start of melatonin feeding and continued to cycle for 1 year or more (N = 6). Plasma prolactin concentrations remained suppressed (less than 20 ng/ml) for the duration of melatonin-feeding (Groups CL/M and NL/M) whereas control values (Group C) were elevated (20-120 ng/ml) between April and August (P less than 0.05). The ovarian response by hinds to melatonin therefore depends on initial reproductive status and recent photoperiodic history, and continued administration to cyclic hinds stimulates prolonged ovarian cyclicity irrespective of the time of year.     

Induction of early breeding in red deer (Cervus elaphus) by melatonin

Lactating (N = 12) and non-lactating (N = 6) red deer hinds and one stag at pasture were given concentrates (500 g/head) containing melatonin (5 mg/head) daily at 16:00 h from 18 June to 16 October. The stag shed the antler velvet and started rutting 5 weeks ahead of untreated stags, and hinds had their first oestrus and ovulation of the breeding season in mid-September, 5 weeks in advance of control lactating (N = 9) and non-lactating (N = 5) hinds. Treated hinds were allowed to mate with the treated stag and control hinds ran in an adjacent paddock with an untreated stag. All hinds became pregnant during the study with all but 2 melatonin-fed hinds (1 lactating, 1 non-lactating) and 1 control hind (lactating) conceiving at the first oestrus. Melatonin-treated hinds lost slightly more live weight than did controls from June to November, but their suckled calves grew throughout at a rate similar to those of control hinds.     

Oestrous cycles and the breeding season of the Père David's deer hind (Elaphurus davidianus)

Reproductive cycles were studied in a group of tame Père David's deer hinds. The non-pregnant hind is seasonally polyoestrous and, in animals studied over 2 years, the breeding season began in early August (2 August +/- 3.3 days; s.e.m., N = 9) and ended in mid-December (18 December +/- 5.7 days; N = 8) and early January (6 January +/- 3.2 days; N = 11) in consecutive years. During the anoestrous period, plasma progesterone concentrations were low (0.2 +/- 0.01 ng/ml) or non-detectable. There was a small, transient increase in progesterone values before the onset of the first cycle of the breeding season. In daily samples taken during an oestrous cycle in which hinds were mated by a marked vasectomized stag, progesterone concentrations remained low (less than 0.5 ng/ml) for a period of about 6 days around the time of oestrus, showed a significant increase above oestrous levels by Day 4 (Day 0 = day of oestrus) and then continued to increase for 18 +/- 2.8 days to reach mean maximum luteal levels of 3.5 +/- 0.6 ng/ml. The plasma progesterone profiles from a number of animals indicated that marking of the hinds by the vasectomized stag did not occur at each ovulation during the breeding season and therefore an estimate of the cycle length could not be determined by this method. In the following year, detection of oestrus in 5 hinds was based on behavioural observations made in the absence of the stag. A total of 19 oestrous cycles with a mean length of 19.5 +/- 0.6 days was observed.(ABSTRACT TRUNCATED AT 250 WORDS)     

Reproductive seasonality in female Iberian red deer (Cervus elaphus hispanicus)

This study characterized the seasonal pattern of luteal cyclicity in Iberian red deer (n=16), by measuring plasma progesterone concentrations in hinds (female red deer) twice per week from calving (May and June) 1996 until May 1997. In eight of these hinds we also examined plasma prolactin profiles to assess seasonal responses to photoperiod. Plasma progesterone concentration in the 16 hinds studied indicated that the reproductive pattern is seasonal, and lasts for 5.73 +/- 0.27 months. After calving, progesterone levels remained basal (no luteal activity) for several months, except in a hind that lost her calf just after calving, and thus did not have to suckle it. This hind showed two consecutive estrus cycles in the month following calving, which suggests that suckling has an inhibiting effect on the resumption of ovarian activity. These results also showed that as long as the hinds do not become pregnant, they show between 5 and 10 estrus cycles per reproductive season (8.06 +/- 0.35), ranging between 105 and 249 days from onset of the first cycle to end of the last one. Uninterrupted cycling lasted for 3.5-6.4 months (mean, 4.6 +/- 0.24). Cyclic luteal activity was found from October to February in all hinds, with a smaller, but notable proportion in September (56.25%) and March (68.8%), whereas it was negligible in the remaining period. Our results show a reproductive season similar to or longer than that recorded by other authors. Prolactin plasma concentrations showed a yearly trend following that of photoperiod, with peak concentrations from April to July, a decrease in August, minimal concentrations from September to February and a sharp increase in March.     

Genetic influences on reproduction of female red deer (Cervus elaphus) (1) seasonal luteal cyclicity

This study compared the onset and duration of the breeding season of female red deer (Cervus elaphus scoticus) and its hybrids with either wapiti (Cervus elaphus nelsoni) or Père David's (PD) deer (Elaphurus davidianus). In Trial 1 (1995), adult red deer (n=9), F1 hybrid wapiti x red deer (n=6) and maternal backcross hybrid PD deer x red deer (i.e., 14 PD; n=9) were maintained together in the presence of a vasectomised red deer stag for 12 months. They were blood-sampled daily or three times weekly so that concentration profiles of plasma progesterone could be used to identify the initiation, duration and cessation of luteal events. There was clear evidence of luteal cyclicity between April and September, with the transition into breeding associated with an apparent silent ovulation and short-lived corpus luteum (i.e., 6-12 days) in every hind. A significant genotype effect occurred in the mean time to first oestrus (P<0.05), with wapiti hybrids and 14 PD hybrids being 9 and 5 days earlier than red deer. Between six and nine oestrous cycles were exhibited by each hind, with no difference in mean cycle length (19.5-19.6 days) between genotypes (P0.10). The overall length of the breeding season was significantly longer for wapiti hybrids (143 days) than for either red deer (130 days) and 14 PD hybrids (132 days, P<0.05).In Trial 2 (1998), adult red deer (n=5), 14 PD hybrids (n=5) and F(1) PD x red deer hybrid (n=5) hinds were maintained together from mid-February (late anoestrus) to early May, in the presence of a fertile red deer stag from 1 April. Thrice-weekly blood sampling yielded plasma progesterone profiles indicative of the onset of the breeding season. Again, there was a significant genotype effect on the mean time to first oestrus (P<0. 05), with F(1) PD hybrids and 14 PD hybrids being 13 and 5 days earlier than red deer. However, conception dates were influenced by the timing of stag joining, and were not significantly different between genotypes. The results indicate genetic effects on reproductive seasonality. However, seasonality observed for PD x red deer hybrids more closely approximated that of red deer than PD deer.     

Plasma concentrations of progesterone in female red deer (Cervus elaphus) during the breeding season, pregnancy and anoestrus

Concentrations of progesterone in peripheral plasma of red deer hinds were basal (less than 1 ng/ml) during lactation/seasonal anoestrus, but increased abruptly at the onset of the breeding season. Lactating hinds (N = 19) started ovarian cycles 10 days later (P less than 0.01) and conceived 16 days later (P less than 0.001) than did 13 weaned hinds. There was no evidence, from plasma progesterone values, of silent oestrus at the start of the season. Progestagen/PMSG treatment induced early ovulations in 8 anoestrous hinds but fertility was low, only 2 conceiving and giving birth. Pregnant hinds (N = 42) had high plasma concentrations of progesterone (mean 3-5 ng/ml) which declined just before parturition.     

Influence of level of nutrition during late pregnancy on reproductive productivity of red deer (2) Adult hinds gestating wapitixred deer crossbred calves

The present study aimed to relate feed intake of red deer hinds in the later stages of gestating wapitixred deer crossbred foetuses on dam body condition, gestation length, birth weight and calf growth. Multiparous hinds (N=18) conceiving at known dates to either wapiti (n=12) or red deer (n=6) sires were housed in individual pens from days 150-220 of pregnancy, during which time they were offered either ad libitum access to pelletised rations (n=6 crossbred-bearing hinds [HH] and n=6 red deer-bearing hinds [RH]) or a restricted offer (n=6 crossbred-bearing hinds [HL]) set at 70% of the average ad libitum intake of HH hind in the previous week. Hinds were returned to pasture at day 220 and calving was closely monitored. Liveweights, body condition score (BCS), and lactation score (LS) of hinds were recorded weekly from day 130 of pregnancy until calves were weaned at 12 weeks of age. Calves were tagged and weighed at birth, and subsequently weighed at 7 and 12 weeks of age. HH and RH hinds exhibited similar patterns and levels of MEI/kg0.75, which peaked at 7.8 MJME/kg0.75 at day 220. HL hinds peaked at approximately 5 MJME/kg0.75 and showed significantly lower rates of liveweight gain during pregnancy. Interestingly, both crossbred-bearing groups initiated mammary development in advance of the RH hinds. While there were significant effects of foetal genotype on mean gestation length (239 days versus 234 days for crossbred versus red deer) and mean birth weight (14.5 kg versus 10 kg), the nutritional contrast for gestation length of crossbred-bearing hinds (i.e. HH versus HL) was not significant but approached significance for birth weight (14.5 kg versus 11.9 kg; P=0.06). Regression analysis revealed weak relationships between changes in hind liveweight and gestation length (P>0.05) but a significant relationship with birth weight (P<0.05). However, change in hind BCS was significantly related to both gestation length and birth weight. Crossbred calves reared by HH hinds were 30% heavier at 7 and 12 weeks of age than the red deer calves. However, those reared by HL hinds were significantly lighter than their genotype contemporaries and only marginally heavier than the red deer calves. These results generally contrast with the previous studies on red deer hinds gestating red deer foetuses [Asher, G.W., Mulley, R.C., O'Neill, K.T., Scott, I.C., Jopson, N.B., Littlejohn, R. 2004. Influence of level of nutrition during late pregnancy on reproductive productivity of red deer, (1) Adult and primiparous hinds gestating red deer calves. Anim. Reprod. Sci., in press] and indicate that the genetically determined higher growth requirements of crossbred foetuses may override any mechanism of compensatory control of gestation length at the expense of calf birth weight. Furthermore, there were marked carryover effects of late gestational feeding on crossbred calf growth and their dam's BCS that highlight the high nutritional demands of lactation.     

Genetic influences on reproduction of female red deer (Cervus elaphus) (2) seasonal and genetic effects on the superovulatory response to exogenous FSH

This study evaluated the influences of seasons and genotype on the superovulatory response to a standardised oFSH regimen in red deer (Cervus elaphus scoticus) and its hybrids with either wapiti (C.e. nelsoni) or Père David's (PD) deer (Elaphurus davidianus). Adult red deer (n=9), F(1) hybrid wapiti x red deer (n=6), and maternal backcross hybrid PD x red deer (i.e., 14 PD hybrid; n=9) were kept together in the presence of a vasectomised stag for 13 months. At 6 weekly intervals, all hinds received a standardised treatment regimen used routinely to induce a superovulatory response in red deer hinds, with 10 consecutive treatments spanning an entire year. This involved synchronisation with intravaginal progesterone devices and delivery of multiple injections of oFSH (equivalent to 72 units NIH-FSH-S(1)). Laparoscopy to assess ovarian response was performed 6-7 days after the removal of the devices. Both season and genotype had significant effects on ovulation rate (OR) and total follicular stimulation (TFS) (P<0.05). For all the three genotypes, ovarian responses were highest from March to November (breeding season) and lowest in the period from December to January, inclusive. Mean OR for red deer hinds ranged from 3.7 to 1.8 during the breeding season, with no observable trend. All red deer hinds were anovulatory during December and January. A similar pattern occurred for 14 PD hybrids, although mean OR during the breeding seasons were twofold lower than for the red deer. For F(1) wapiti hybrids, the first two treatments in March and April resulted in the highest mean OR observed (15.6 and 11.7, respectively). Thereafter, mean values ranged between 6.3 and 4.7 for the remainder of the breeding season. Furthermore, mean OR of 3.0 and 0.5 were recorded in December and January, respectively. For the red deer and F(1) wapiti hybrids, between-hind variation in OR was not randomly distributed across the treatment dates, indicating that the individuals varied significantly in their ability to respond to oFSH, at least within a given season.In conclusion, the study has shown that relative to red deer, F(1) wapiti hybrid hinds exhibit a higher sensitivity to oFSH, whereas 14 PD hybrid hinds have a lower sensitivity. However, individual variation within genotype was very marked. A seasonal effect was apparent for all genotypes, although some F(1) wapiti hybrid hinds exhibited ovulatory responses throughout the year.     

Immunoreactive luteinizing hormone, estradiol, progesterone, testosterone, and androstenedione levels during the breeding season and anestrus in Siberian tigers

Seasonal analysis of 1239 captive births of Siberian tigers (Panthera tigris altaica) indicated a peak in April to June (P less than 0.001). Studies on seven animals in Minnesota indicated that behavioral heat cycles and ovarian follicular phase cycles began in late January and ceased in early June. Behavioral observation of 12 heat cycles in four tigers yielded an estrous length of 5.3 +/- 0.2 days and an interestrous interval of 25.0 +/- 1.3 days. Hormone assays on weekly blood samples (N = 180) from three female tigers indicated 16 cycles in two breeding seasons. Peak estradiol-17 beta levels were 46.7 +/- 6.0 pg/ml (N = 17) and interestrous concentrations were 8.7 +/- 0.66 pg/ml (N = 28) during the breeding season. Anestrous estradiol levels were 4.2 +/- 0.5 pg/ml (N = 70). The interestrous interval between estradiol peaks was 24.9 +/- 1.3 days (N = 9) with two outliers of 42 days. Serum progesterone concentrations from February to June were 1.2 +/- 0.15 ng/ml (N = 32), providing no evidence for ovulation or corpus luteum formation. Luteinizing hormone (LH) levels were 0.56 +/- 0.04 ng/ml (N = 180). Serum testosterone (r=0.71, P less than 0.001) and androstenedione levels (r=0.75, P less than 0.001) were correlated with estradiol during the breeding season. The duration of anestrus was 8 mo in two of these tigers. The interval was shortened in one tiger by exposure to a 16L:8D photoperiod. The Siberian tiger appears to be a polyestrous seasonal breeder and an induced ovulator whose breeding season may be synchronized by photoperiod.     

Response in peripheral plasma melatonin to photoperiod change and the effects of exogenous melatonin on seasonal quiescence in the tammar, Macropus eugenii

During the light phase of each of 3 photoperiods tested, plasma melatonin concentrations were less than 16 to 62 pg/ml and during the dark phase they were 31 to 169 pg/ml. When the photoperiod to which the tammars were exposed was altered from 15 h light:9 h dark to 12L:12D the onset of the nocturnal rise in melatonin was advanced from the first day, thereby extending its duration, and the females gave birth 32 +/- 0.4 (mean +/- s.e.m.) days later. To test whether melatonin mediated this effect of photo-period change, tammars in a second group were injected s.c. with melatonin (400 ng/kg, N = 6) or the arachis oil vehicle (N = 6), 2.5 to 2.25 h before dark during 15L:9D for 15 days before exposure to 12.5L:11.5D. The melatonin injections mimicked the endogenous melatonin profile of 12L:12D and the melatonin-injected tammars gave birth 32 +/- 0.8 days after the start of injections, which was the same as the interval from photoperiod change in Exp. 1 but was significantly different (P less than 0.005) from the interval in the control group (46.0 +/- 1.1 days). These results show that exogenous melatonin given 2.5 to 2.25 h in advance of the endogenous rise fully mimics the response of the tammar to photoperiod change.     

Efficacy of intermittent or continuous administration of GnRH in inducing ovulation in early and late seasonal anoestrus in the Père David's deer hind (Elaphurus davidianus)

Père David's deer hinds were treated with GnRH, administered as intermittent i.v. injections (2.0 micrograms/injection at 2-h intervals) for 4 days, or as a continuous s.c. infusion (1.0 micrograms/h) for 14 days. These treatments were given early (February-March) and late (May-June) in the period of seasonal anoestrus. The administration of repeated injections of GnRH increased mean LH concentrations from pretreatment values of 0.54 +/- 0.09 to 2.10 +/- 0.25 ng/ml over the first 8 h of treatment in early anoestrus, and from 0.62 +/- 0.11 to 2.73 +/- 0.49 ng/ml in late anoestrus. The mean amplitude of GnRH-induced LH episodes was greater (P less than 0.01) in late (4.03 +/- 0.28 ng/ml) than in early (3.12 +/- 0.26 ng/ml) anoestrus, but within each replicate (early or late anoestrus), neither mean LH episode amplitude nor mean plasma LH concentrations differed significantly between the four periods of intensive blood sampling. On the basis of their progesterone profiles, 6/12 hinds had ovulated in response to treatment with injections of GnRH (1/6 in early anoestrus and 5/6 in late anoestrus), and oestrus and a preovulatory LH surge were recorded in all of these animals. Oestrus and a preovulatory LH surge were also recorded in one other animal treated in early anoestrus in which progesterone concentrations remained low. The mean times of onset of oestrus (91.0 +/- 1.00 and 62.4 +/- 0.98 h) and of the preovulatory LH surge (85.8 +/- 3.76 and 59.4 +/- 0.25 h) both occurred significantly earlier (P less than 0.001) in animals treated in late anoestrus. Continuous infusion of GnRH to seasonally anoestrous hinds resulted in an increase in mean plasma LH concentrations, but this response did not differ significantly between early (2.15 +/- 0.28 ng/ml) and late (2.48 +/- 0.26 ng/ml) anoestrus. Ovulation, based on progesterone profiles, occurred in 2/7 hinds in early anoestrus and in 4/6 hinds in late anoestrus. Oestrus was detected in all except one of these hinds. The mean time of onset of oestrus occurred earlier in animals treated in late anoestrus (66.2 +/- 0.32 h and 46.7 +/- 0.67 h, P less than 0.01). The administration of GnRH, given either intermittently or continuously, will induce ovulation in a proportion of seasonally anoestrous Père David's deer. However, more animals ovulate in response to this treatment in late than in early anoestrus (75% compared with 23%).     

Seasonal and daily variations in plasma melatonin in the high-arctic Svalbard ptarmigan (Lagopus mutus hyperboreus).

This study presents the daily rhythm of melatonin secretion throughout one year in a bird from the northern hemisphere, the Svalbard ptarmigan (Lagopus mutus hyperboreus), which lives naturally at 76-80 degrees N. Eight Svalbard ptarmigan were caged outdoors at 70 degrees N and blood sampled throughout one day each month for 13 months. At this latitude, daylight is continuous between May and August, but there is a short period of civil twilight around noon from late November to mid January. There was no daily rhythm in plasma melatonin in May-July. Plasma melatonin levels varied significantly throughout the day in all other months of the year, with the nighttime increase reflecting the duration of darkness. The highest mean plasma concentration occurred at midnight in March (110.1 +/- 16.5 pg/ml) and represented the annual peak in estimated daily production. Around the winter solstice, melatonin levels were significantly reduced at noon and elevated during the nearly 18 h of consecutive darkness, and the estimated mean daily production of melatonin was significantly reduced. Thus, at the times of the year characterized by light-dark cycles, melatonin may convey information concerning the length of the day and, therefore, progression of season. The nearly undetectable low melatonin secretion in summer and the reduced amplitude and production in midwinter indicate a flexible circadian system that may reflect an important adaptation to life in the Arctic.     

Influence of prenatal photoperiod on postnatal prolactin secretion in red deer (Cervus elaphus).

Plasma prolactin concentrations were higher (P < 0.001) in newborn red deer calves whose mothers had been maintained for the last 14 weeks of gestation in long days (18 h light) (group L, n = 9) than in those whose mothers had been kept over the same period in short days (6 h light) (group S, n = 5). After transfer of all hinds and suckled calves on the day of birth to constant intermediate daylength (12 h light), prolactin concentrations decreased exponentially (P < 0.001) in group L calves, but not in group S, during the first 21 days. Thereafter, prolactin fell to a nadir in group L calves and rose to peak values in group S calves at 8-12 weeks post partum (P = 0.003), before converging again by 14 weeks. The pattern of prolactin secretion over the first 14 weeks of life was therefore significantly affected by prenatal photoperiod. Plasma prolactin concentrations in the adult hinds were higher (P < 0.001) in group L than group S at 4-10 weeks before parturition; they were similarly high around parturition and fell thereafter to baseline values after 7 weeks. These results provide evidence that deer fetuses respond to photoperiodic information, thereby acquiring a photoperiodic history in utero that influences postnatal responses to photoperiod.     

The influence of birth date on the development of seasonal cycles in red deer hinds (Cervus elaphus)

The relative importance of age and photoperiod in the timing of seasonal cycles was studied in red deer during the first 18 months of life. Seasonal cycles of appetite, live-weight change, coat growth, oestrus and metabolic rate were examined in two groups of seven hind calves born in early May (group E) or nine weeks later in July (group L).
There was no difference between groups in the timing or amplitude of the seasonal voluntary food intake (VFI) cycle, or in the timing of seasonal changes in rate of live-weight gain, although mean live-weight gain was significantly greater for group L than group E during the first winter of life.
Four animals in group L retained their calf coats throughout the first winter. In group L, the first summer coat fibres appeared approximately four weeks earlier, and mean fibre length for the subsequent winter coat was longer in group L than in group E. There was no difference between groups in the timing or amplitude of the seasonal cycle of plasma prolactin concentration, or in the incidence and mean time of onset of puberty.
There was no difference between groups in mean energy expenditure per unit of live weight at any time of the year. Mean daily energy expenditure was significantly higher in summer than in winter, but values measured when the animals were resting and corrected for differences in VFI were similar throughout the year.
It is concluded that the seasonal cycles of appetite, live-weight change and oestrus in young red deer hinds are not modified by age, although the normal cycle of coat replacement can be disturbed in animals born towards the end of the calving season.     

Production of interferon by red deer (Cervus elaphus) conceptuses and the effects of roIFN-tau on the timing of luteolysis and the success of asynchronous embryo transfer

The role of interferon in early pregnancy in red deer was investigated by (a) measuring production of interferon by the conceptus, (b) testing the anti-luteolytic effect of recombinant interferon-tau in non-pregnant hinds, and (c) treatment of hinds with interferon after asynchronous embryo transfer. Blastocysts were collected from 34 hinds by uterine flushing 14 (n = 2), 16 (n = 2), 18 (n = 8), 20 (n = 13) or 22 (n = 9) days after synchronization of oestrus with progesterone withdrawal. Interferon anti-viral activity was detectable in uterine flushings from day 16 to day 22, and increased with duration of gestation (P < 0.01) and developmental stage (P < 0.01). When interferon-tau was administered daily between day 14 and day 20 to non-pregnant hinds to mimic natural blastocyst production, luteolysis was delayed by a dose of 0.2 mg day(-1) (27.3 +/- 1.3 days after synchronization, n = 4 versus 21 +/- 0 days in control hinds, n = 3; P < 0.05). Interferon-tau was administered to hinds after asynchronous embryo transfer to determine whether it protects the conceptus against early pregnancy loss. Embryos (n = 24) collected on day 6 from naturally mated, superovulated donors (n = 15) were transferred into synchronized recipients on day 10 or day 11. Interferon-tau treatment (0.2 mg daily from day 14 to 20) increased calving rate from 0 to 64% in all recipients (0/11 versus 7/11, P < 0.005), and from 0 to 67% in day 10 recipients (0/8 versus 6/9, P < 0.01). The increased success rate of asynchronous embryo transfer after interferon-tau treatment in cervids may be of benefit where mismatched embryo-maternal signalling leads to failure in the establishment of pregnancy.     

Annual cycle of liveweight and reproductive changes of farmed male fallow deer (Dama dama) and the effect of daily oral administration of melatonin in summer on the attainment of seasonal fertility

Entire bucks (N = 7) exhibited pronounced liveweight gains over spring and summer months (October-February), to reach a peak mean weight of 59.8 kg, and rapid liveweight losses over the rutting period (April-May) with a minimum mean liveweight of 54.2 kg. Mean neck girth and serum testosterone levels increased during late summer (January-March) and peaked at 387 mm and 12 ng/ml respectively immediately before the onset of the rut (April). Thereafter both measures declined during winter and spring months (June-December). Bucks castrated prepubertally (N = 11) exhibited similar but less pronounced changes in mean liveweight and neck girth, in the absence of any change in testosterone secretion. Peak mean testicular diameter of entire bucks (39 mm) occurred immediately before the rut and was followed by testicular regression over winter and spring months (June-November), such that the testes attained their minimum mean size of 18 mm diameter in early summer (November). Motile spermatozoa were absent from ejaculates collected in summer (November 1983, 1984; January 1984). However, ejaculates collected pre-rut (late March), immediately post-rut (June) and in early spring (September) contained successively increasing numbers of motile spermatozoa. A further 14 polled, entire bucks were given orally 5 mg (N = 7; Group A) or 20 mg (N = 7; Group B) melatonin at 15:30 h daily from 1 December 1983 to 14 January 1984 (45 days). Seven control bucks (Group C) received vehicle ration only. The measurements taken for bucks in Groups A and B were not significantly different (P greater than 0.1) on any sampling date and the data for these 2 groups were pooled. Mean serum testosterone concentrations and mean ejaculate volume were not significantly different between melatonin-treated and control bucks on any sampling date, although other measures exhibited significant differences (P less than 0.05) at various treatment or post-treatment dates: melatonin-treated bucks showed a transiently greater increase in neck muscle development during and immediately after treatment, a slight retardation of liveweight gain between 45 and 75 days after treatment, an earlier peak in maximum mean testicular diameter and an earlier onset of sperm presence in ejaculates.     

A comparison of the calving behaviour of farmed adult and yearling red deer (Cervus elaphus) hinds

Mortality of newborn red deer (Cervus elaphus) calves is a major concern on New Zealand farms, as perinatal losses average approximately 10% of calves born. Primiparous red deer (calving as yearlings) lose more calves than multiparous hinds (adults). We performed a study on yearling and adult red deer hinds in order to improve knowledge of their calving behaviour and determine any apparent reasons for calf mortality. Pacing along fence lines was observed frequently during the 24 h period before birth (individuals were pacing in 43% of observations during this period). Adult hinds had a significantly earlier onset and longer duration of pacing (P<0.05). Hinds were often observed isolated (>20 m) from the rest of the herd during the 2 days prior to parturition, and this behaviour was also observed earlier in the adults than in the yearlings (P<0.05). Forty-four percent of adult hinds and 60% of yearlings experienced some form of interference from other hinds during parturition. Among hinds that were interfered with, yearlings had a higher average number of interferences from other hinds during parturition than adults (P<0.05). Calves born to yearling hinds took significantly longer to suckle for the first time (mean=44 min) from their dams than calves born to adult hinds (33 min; P<0.05). We concluded that an inability of hinds to express natural isolation behaviour was likely to contribute to calf mortality due to increased anxiety (indicated by fence line pacing) and encroachment on the birth areas of others, and that calves of yearlings were at an increased risk of mortality due to the higher number of interferences taking place during parturition and the longer time interval between birth and suckling.     

Seasonal luteal cyclicity of pubertal and adult red deer (Cervus elaphus)

Reproductive failure of rising-two-year-old (R(2)) hinds and seasonal misalignment between calving and pastoral feed production are two factors limiting reproductive productivity of farmed red deer hinds in New Zealand. This study aimed to better understand processes around female puberty and breeding seasonality by describing the potential breeding season (i.e., oestrous cyclicity) of three red deer genotypes. A total of 27 hinds born in December 2005, representing Eastern European (Cervus elaphus hippelaphus), Western European (C.e. scoticus) and F1 crossbred (C.e. hippelaphus×scoticus) red deer, were blood sampled thrice-weekly for 7-8 months (February-September/October) across two years spanning the potential breeding seasons as R(2)'s in 2007 (i.e., puberty) and as adults in 2008. Plasma progesterone profiles were used to construct breeding cycle histories for each hind. Four R(2) hinds failed to initiate oestrous cycles (i.e., puberty failure). The remaining R(2) hinds, including all F1 hinds, exhibited between two and seven oestrous cycles. F1 hinds were significantly earlier to initiate, and later to terminate, cyclic activity, resulting in a longer mean pubertal breeding season (139 days) than for Eastern (86 days) and Western hinds (86 days). However, the data for R(2) hinds are confounded by live-weight, with the F1 hinds being significantly heavier than other genotypes. There were significant correlations between live-weight and seasonality parameters in 2007. All hinds were cyclic as adults in 2008, exhibiting between four and nine oestrous cycles, and a mean breeding season duration of between 132 (Western) and 137 (F1) days. For adult hinds there were no significant genotype differences in cyclic onset and cessation timing, and no observable relationships between live-weight and any reproductive parameter. However, the mean dates for the onset of the breeding season for all genotypes in 2008 were 2-3 weeks later than normally expected for adult hinds in New Zealand. The reasons for this are unclear but may relate to chronic stress of frequent animal handling. The study has demonstrated that puberty in red deer hinds is associated with a shorter potential breeding season than for adult hinds, and that perturbation of breeding activity appears to be quite common, leading to incidences of puberty failure and possibly other aberrant cyclic events. Live-weight×genotype interactions may influence puberty but do not appear to be strongly expressed in adults. However, the relatively late onset of oestrous cyclicity in the adult hinds may be an artefact of the study that has masked genetic influences on seasonal breeding patterns.     

Reproductive refractoriness in the Welsh Mountain ewe induced by a short photoperiod can be overridden by exposure to a shorter photoperiod

The objectives were to determine if relative lengths of photoperiods that induce reproductive cycles in ewes affect the length of the subsequent breeding season, if duration of the refractoriness that terminates breeding is affected by photoperiod length, and if the resulting refractoriness to an inductive photoperiod is absolute. Groups of Welsh Mountain ewes were exposed to either 12L:12D (n = 12) or 8L:16D (n = 6) photoperiods beginning at the summer solstice when daylengths reach a maximum of 17.5 h at Bristol, England. A control group (n = 10) was exposed to natural daylengths. Ovarian cycles in the controls, as judged by monitored plasma progesterone levels, commenced in early October, about 1 mo later (p less than 0.001 in both cases) than in sheep exposed to 12L:12D or 8L:16D. The advancement in cycle onset was similar under 12L:12D and 8L:16D (69 +/- 2 and 77 +/- 4 days after the summer solstice compared with 102 +/- 2 days in the controls). Duration of the breeding season (100 +/- 4 days) in ewes exposed to 12L:12D was significantly shorter (p less than 0.001 in both cases) than in ewes exposed to natural daylengths or 8L:16D (153 +/- 3 and 133 +/- 5 days, respectively). Approximately 70 days after the ending of ovulatory cycles in the 12L:12D group, half of the animals (n = 6) were transferred to 8L:16D. This treatment greatly (p less than 0.001) reduced the duration of anestrus and cycles began again 62 +/- 4 days after transfer to 8L:16D, or about 90 days earlier than in ewes (n = 6) remaining in 12L:12D.(ABSTRACT TRUNCATED AT 250 WORDS)     

Ontogeny of the daily profile of plasma melatonin in European starlings raised under long or short photoperiods

Photoperiodic manipulation of young European starlings suggests that their reproductive physiology is incapable of responding to a short photoperiod until they are fully grown. This study aimed to determine whether the lack of response to a short photoperiod is reflected in the daily profile of plasma melatonin concentrations. Five-day-old starlings taken from nest boxes showed a significant (p < 0.0001) rhythm in plasma melatonin concentrations, with high values during night. In nestlings hand-reared from 5 days of age on a long photoperiod (LD 16:8), equivalent to natural photoperiod at the time, the amplitude of the daily rhythm in melatonin increased significantly (p < 0.01) with age until birds were fully grown (20 days old). In nestlings reared on a short photoperiod (LD 8:16), the daily melatonin profile remained almost identical to that of long photoperiod birds until they were fully grown. However, after 20 days old, the duration of elevated nighttime melatonin began to extend to encompass the entire period of darkness. In contrast, fully grown starlings transferred from a long to a short photoperiod had partially adapted to the short photoperiod after 5 days; by 10 days, the daily melatonin profile was identical to that of birds held chronically on a short photoperiod. Thus, consistent with responses of reproductive physiology, the pineal of young birds appears to be incapable of perceiving, or adapting to, a short photoperiod.